{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,2]],"date-time":"2026-11-02T11:29:37Z","timestamp":1793618977784,"version":"4.5.0"},"reference-count":89,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2025,5,1]],"date-time":"2025-05-01T00:00:00Z","timestamp":1746057600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"},{"start":{"date-parts":[[2025,5,1]],"date-time":"2025-05-01T00:00:00Z","timestamp":1746057600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"}],"funder":[{"DOI":"10.13039\/501100021856","name":"Ministero dell\u2019Universit\u00e0 e della Ricerca","doi-asserted-by":"publisher","award":["PRIN 2022-NAZ-0168"],"award-info":[{"award-number":["PRIN 2022-NAZ-0168"]}],"id":[{"id":"10.13039\/501100021856","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100003196","name":"Ministero della Salute","doi-asserted-by":"publisher","award":["Ricerca Corrente"],"award-info":[{"award-number":["Ricerca Corrente"]}],"id":[{"id":"10.13039\/501100003196","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"DOI":"10.1038\/s41598-025-98595-8","type":"journal-article","created":{"date-parts":[[2025,5,1]],"date-time":"2025-05-01T16:13:32Z","timestamp":1746116012000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":7,"title":["Visuomotor paired associative stimulation enhances corticospinal excitability in post-stroke patients with upper-limb hemiparesis"],"prefix":"10.1038","volume":"15","author":[{"given":"Michela","family":"Picardi","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Giacomo","family":"Guidali","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Antonio","family":"Caronni","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Viviana","family":"Rota","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Massimo","family":"Corbo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nadia","family":"Bolognini","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2025,5,1]]},"reference":[{"key":"98595_CR1","doi-asserted-by":"publisher","first-page":"1035","DOI":"10.1053\/apmr.2002.33984","volume":"83","author":"NE Mayo","year":"2002","unstructured":"Mayo, N. E., Wood-Dauphinee, S., C\u00f4t\u00e9, R., Durcan, L. & Carlton, J. Activity, participation, and quality of life 6 months poststroke. Arch. Phys. Med. Rehabil. 83, 1035\u20131042 (2002).","journal-title":"Arch. Phys. Med. Rehabil"},{"key":"98595_CR2","doi-asserted-by":"publisher","first-page":"342","DOI":"10.1002\/ana.24734","volume":"80","author":"CE Lang","year":"2016","unstructured":"Lang, C. E. et al. Dose response of task-specific upper limb training in people at least 6 months poststroke: a phase II, single-blind, randomized, controlled trial. Ann. Neurol. 80, 342\u2013354 (2016).","journal-title":"Ann. Neurol."},{"key":"98595_CR3","first-page":"CD010820","volume":"11","author":"A Pollock","year":"2014","unstructured":"Pollock, A. et al. Interventions for improving upper limb function after stroke (Cochrane review) [with consumer summary]. Cochrane Database Syst. Reviews. 11, CD010820\u2013CD010820 (2014).","journal-title":"Cochrane Database Syst. Reviews"},{"key":"98595_CR4","doi-asserted-by":"publisher","unstructured":"Su, F. & Xu, W. Enhancing brain plasticity to promote stroke recovery. Front. Neurol. 2020, 11. https:\/\/doi.org\/10.3389\/fneur.2020.554089  (2020).","DOI":"10.3389\/fneur.2020.554089"},{"key":"98595_CR5","doi-asserted-by":"publisher","first-page":"953","DOI":"10.1179\/1743132811Y.0000000045","volume":"33","author":"MA Ahmed","year":"2011","unstructured":"Ahmed, M. A., Mohamed, S. A. & Sayed, D. Long-term antalgic effects of repetitive transcranial magnetic stimulation of motor cortex and serum beta-endorphin in patients with Phantom pain. Neurol. Res. 33, 953\u2013958 (2011).","journal-title":"Neurol. Res."},{"key":"98595_CR6","doi-asserted-by":"publisher","first-page":"190","DOI":"10.3390\/healthcare10020190","volume":"10","author":"S Anwer","year":"2022","unstructured":"Anwer, S. et al. Rehabilitation of upper limb motor impairment in stroke: a narrative review on the prevalence, risk factors, and economic statistics of stroke and state of the Art therapies. Healthcare 10, 190 (2022).","journal-title":"Healthcare"},{"key":"98595_CR7","doi-asserted-by":"publisher","unstructured":"Di Pino, G. et al. Modulation of brain plasticity in stroke: a novel model for neurorehabilitation. Nat. Rev. Neurol.   10, 597\u2013608. https:\/\/doi.org\/10.1038\/nrneurol.2014.162   (2014).","DOI":"10.1038\/nrneurol.2014.162"},{"key":"98595_CR8","doi-asserted-by":"publisher","first-page":"113484","DOI":"10.1016\/j.bbr.2021.113484","volume":"414","author":"G Guidali","year":"2021","unstructured":"Guidali, G., Roncoroni, C. & Bolognini, N. Paired associative stimulations: novel tools for interacting with sensory and motor cortical plasticity. Behav. Brain. Res. 414, 113484 (2021).","journal-title":"Behav. Brain. Res."},{"key":"98595_CR9","doi-asserted-by":"publisher","first-page":"658723","DOI":"10.3389\/fnhum.2021.658723","volume":"15","author":"G Guidali","year":"2021","unstructured":"Guidali, G., Roncoroni, C. & Bolognini, N. Modulating frontal networks\u2019 Timing-Dependent-Like plasticity with paired associative stimulation protocols: recent advances and future perspectives. Front. Hum. Neurosci. 15, 658723 (2021).","journal-title":"Front. Hum. Neurosci."},{"key":"98595_CR10","doi-asserted-by":"publisher","first-page":"2140","DOI":"10.1016\/j.clinph.2017.08.003","volume":"128","author":"A Suppa","year":"2017","unstructured":"Suppa, A. et al. The associative brain at work: evidence from paired associative stimulation studies in humans. Clin. Neurophysiol. 128, 2140\u20132164 (2017).","journal-title":"Clin. Neurophysiol."},{"key":"98595_CR11","doi-asserted-by":"publisher","first-page":"572","DOI":"10.1093\/brain\/123.3.572","volume":"123","author":"K Stefan","year":"2000","unstructured":"Stefan, K., Kunesch, E., Cohen, L. G., Benecke, R. & Classen, J. Induction of plasticity in the human motor cortex by paired associative stimulation. Brain 123, 572\u2013584 (2000).","journal-title":"Brain"},{"key":"98595_CR12","doi-asserted-by":"publisher","first-page":"2339","DOI":"10.1152\/jn.00900.2002","volume":"89","author":"A Wolters","year":"2003","unstructured":"Wolters, A. et al. A temporally asymmetric hebbian rule governing plasticity in the human motor cortex. J. Neurophysiol. 89, 2339\u20132345 (2003).","journal-title":"J. Neurophysiol."},{"key":"98595_CR13","doi-asserted-by":"publisher","first-page":"567","DOI":"10.3390\/neurolint16030043","volume":"16","author":"A Baroni","year":"2024","unstructured":"Baroni, A. et al. The effectiveness of paired associative stimulation on motor recovery after stroke: a scoping review. Neurol. Int. 16, 567\u2013589 (2024).","journal-title":"Neurol. Int."},{"key":"98595_CR14","doi-asserted-by":"publisher","first-page":"732","DOI":"10.1016\/j.clinph.2015.04.072","volume":"127","author":"M Wischnewski","year":"2016","unstructured":"Wischnewski, M. & Schutter, D. J. L. G. Efficacy and time course of paired associative stimulation in cortical plasticity: implications for neuropsychiatry. Clin. Neurophysiol. 127, 732\u2013739 (2016).","journal-title":"Clin. Neurophysiol."},{"key":"98595_CR15","doi-asserted-by":"publisher","first-page":"1","DOI":"10.3389\/fnins.2019.00792","volume":"13","author":"J Silverstein","year":"2019","unstructured":"Silverstein, J. et al. Paired associative stimulation as a tool to assess plasticity enhancers in chronic stroke. Front. Neurosci. 13, 1\u201310 (2019).","journal-title":"Front. Neurosci."},{"key":"98595_CR16","doi-asserted-by":"publisher","first-page":"671","DOI":"10.1177\/1545968318785043","volume":"32","author":"JK Ferris","year":"2018","unstructured":"Ferris, J. K., Neva, J. L., Francisco, B. A. & Boyd, L. A. Bilateral motor cortex plasticity in individuals with chronic stroke, induced by paired associative stimulation. Neurorehabil. Neural Repair. 32, 671\u2013681 (2018).","journal-title":"Neurorehabil. Neural Repair."},{"key":"98595_CR17","doi-asserted-by":"publisher","first-page":"627","DOI":"10.1016\/j.brs.2020.01.017","volume":"13","author":"G Guidali","year":"2020","unstructured":"Guidali, G., Carneiro, M. I. S. & Bolognini, N. Paired associative stimulation drives the emergence of motor resonance. Brain Stimul. 13, 627\u2013636 (2020).","journal-title":"Brain Stimul"},{"key":"98595_CR18","doi-asserted-by":"publisher","first-page":"102128","DOI":"10.1016\/j.pneurobio.2021.102128","volume":"205","author":"D Kemmerer","year":"2021","unstructured":"Kemmerer, D. What modulates the mirror neuron system during action observation? Multiple factors involving the action, the actor, the observer, the relationship between actor and observer, and the context. Prog Neurobiol. 205, 102128 (2021).","journal-title":"Prog Neurobiol."},{"key":"98595_CR19","doi-asserted-by":"publisher","first-page":"169","DOI":"10.1146\/annurev.neuro.27.070203.144230","volume":"27","author":"G Rizzolatti","year":"2004","unstructured":"Rizzolatti, G. & Craighero, L. The Mirror-Neuron system. Annu. Rev. Neurosci. 27, 169\u2013192 (2004).","journal-title":"Annu. Rev. Neurosci."},{"key":"98595_CR20","doi-asserted-by":"publisher","first-page":"139","DOI":"10.1016\/j.cortex.2023.03.006","volume":"163","author":"G Guidali","year":"2023","unstructured":"Guidali, G., Picardi, M., Gramegna, C. & Bolognini, N. Modulating motor resonance with paired associative stimulation: neurophysiological and behavioral outcomes. Cortex 163, 139\u2013153 (2023).","journal-title":"Cortex"},{"key":"98595_CR21","doi-asserted-by":"publisher","first-page":"121082","DOI":"10.1016\/j.neuroimage.2025.121082","volume":"308","author":"G Guidali","year":"2025","unstructured":"Guidali, G., Arrigoni, E., Bolognini, N. & Pisoni, A. M1 large-scale network dynamics support human motor resonance and its plastic reshaping. Neuroimage 308, 121082 (2025).","journal-title":"Neuroimage"},{"key":"98595_CR22","doi-asserted-by":"publisher","first-page":"257","DOI":"10.3390\/brainsci15030257","volume":"15","author":"G Guidali","year":"2025","unstructured":"Guidali, G. & Bolognini, N. Tracking changes in corticospinal excitability during visuomotor paired associative stimulation to predict motor resonance rewriting. Brain Sci. 15, 257 (2025).","journal-title":"Brain Sci."},{"key":"98595_CR23","doi-asserted-by":"publisher","unstructured":"Thieme, H. et al. Mirror therapy for improving motor function after stroke. Cochrane Database Syst. Rev.  2018, 45.  https:\/\/doi.org\/10.1002\/14651858.CD008449.pub3   (2018).","DOI":"10.1002\/14651858.CD008449.pub3"},{"key":"98595_CR24","doi-asserted-by":"publisher","unstructured":"Zhang, C., Li, X. & Wang, H. Application of action observation therapy in stroke rehabilitation: a systematic review. Brain Behav. 13, 56.  https:\/\/doi.org\/10.1002\/brb3.3157 (2023).","DOI":"10.1002\/brb3.3157"},{"key":"98595_CR25","doi-asserted-by":"publisher","first-page":"113","DOI":"10.1007\/s00221-006-0844-5","volume":"180","author":"E Castel-Lacanal","year":"2007","unstructured":"Castel-Lacanal, E., Gerdelat-Mas, A., Marque, P., Loubinoux, I. & Simonetta-Moreau, M. Induction of cortical plastic changes in wrist muscles by paired associative stimulation in healthy subjects and post-stroke patients. Exp. Brain Res. 180, 113\u2013122 (2007).","journal-title":"Exp. Brain Res."},{"key":"98595_CR26","doi-asserted-by":"publisher","first-page":"366","DOI":"10.1177\/1545968308322841","volume":"825","author":"E Castel-Lacanal","year":"2009","unstructured":"Castel-Lacanal, E., Marque, P. & Tardy, J. Induction of cortical plastic changes in wrist muscles by paired associative stimulation in the recovery phase of stroke patients. Neurorehabil Neural Repair. 825, 366\u2013372 (2009).","journal-title":"Neurorehabil Neural Repair."},{"key":"98595_CR27","doi-asserted-by":"publisher","first-page":"2608","DOI":"10.1152\/jn.1995.73.6.2608","volume":"73","author":"L Fadiga","year":"1995","unstructured":"Fadiga, L., Fogassi, L., Pavesi, G. & Rizzolatti, G. Motor facilitation during action observation: a magnetic stimulation study. J. Neurophysiol. 73, 2608\u20132611 (1995).","journal-title":"J. Neurophysiol."},{"key":"98595_CR28","doi-asserted-by":"publisher","first-page":"15933","DOI":"10.1038\/s41598-023-43227-2","volume":"13","author":"G Guidali","year":"2023","unstructured":"Guidali, G., Picardi, M., Franca, M., Caronni, A. & Bolognini, N. The social relevance and the Temporal constraints of motor resonance in humans. Sci. Rep. 13, 15933 (2023).","journal-title":"Sci. Rep."},{"key":"98595_CR29","doi-asserted-by":"publisher","first-page":"331","DOI":"10.1016\/j.neuropsychologia.2014.09.034","volume":"64","author":"KR Naish","year":"2014","unstructured":"Naish, K. R., Houston-Price, C., Bremner, A. J. & Holmes, N. P. Effects of action observation on corticospinal excitability: muscle specificity, direction, and timing of the mirror response. Neuropsychologia 64, 331\u2013348 (2014).","journal-title":"Neuropsychologia"},{"key":"98595_CR30","doi-asserted-by":"publisher","first-page":"269","DOI":"10.1016\/j.clinph.2020.10.003","volume":"132","author":"S Rossi","year":"2021","unstructured":"Rossi, S. et al. Safety and recommendations for TMS use in healthy subjects and patient populations, with updates on training, ethical and regulatory issues: expert guidelines. Clin. Neurophysiol. 132, 269\u2013306 (2021).","journal-title":"Clin. Neurophysiol."},{"key":"98595_CR31","doi-asserted-by":"publisher","first-page":"1149","DOI":"10.3758\/BRM.41.4.1149","volume":"41","author":"F Faul","year":"2009","unstructured":"Faul, F., Erdfelder, E., Buchner, A. & Lang, A. G. Statistical power analyses using G*Power 3.1: tests for correlation and regression analyses. Behav. Res. Methods. 41, 1149\u20131160 (2009).","journal-title":"Behav. Res. Methods"},{"key":"98595_CR32","doi-asserted-by":"publisher","first-page":"13","DOI":"10.1016\/S1567-424X(09)70205-3","volume":"56","author":"F Awiszus","year":"2003","unstructured":"Awiszus, F. TMS and threshold hunting. Suppl. Clin. Neurophysiol. 56, 13\u201323 (2003).","journal-title":"Suppl. Clin. Neurophysiol."},{"key":"98595_CR33","doi-asserted-by":"publisher","first-page":"1758","DOI":"10.1002\/jnr.24319","volume":"96","author":"T Dissanayaka","year":"2018","unstructured":"Dissanayaka, T., Zoghi, M., Farrell, M., Egan, G. & Jaberzadeh, S. Comparison of Rossini\u2013Rothwell and adaptive threshold-hunting methods on the stability of TMS induced motor evoked potentials amplitudes. J. Neurosci. Res. 96, 1758\u20131765 (2018).","journal-title":"J. Neurosci. Res."},{"key":"98595_CR34","doi-asserted-by":"publisher","first-page":"456","DOI":"10.1177\/1545968311427406","volume":"26","author":"M Franceschini","year":"2012","unstructured":"Franceschini, M. et al. Clinical relevance of action observation in upper-limb stroke rehabilitation: a possible role in recovery of functional dexterity. A randomized clinical trial. Neurorehabil Neural Repair. 26, 456\u2013462 (2012).","journal-title":"Neurorehabil Neural Repair."},{"key":"98595_CR35","doi-asserted-by":"crossref","unstructured":"Tropea, P. et al. Action observation treatment for upper limb rehabilitation in patients with stroke: protocol for a multicenter randomized controlled trial. JMIR Res. Protoc. 12, 45 (2023).","DOI":"10.2196\/42094"},{"key":"98595_CR36","doi-asserted-by":"publisher","first-page":"78","DOI":"10.1016\/j.rehab.2017.11.002","volume":"61","author":"M Tarri","year":"2018","unstructured":"Tarri, M. et al. Five-day course of paired associative stimulation fails to improve motor function in stroke patients. Ann. Phys. Rehabil Med. 61, 78\u201384 (2018).","journal-title":"Ann. Phys. Rehabil Med."},{"key":"98595_CR37","doi-asserted-by":"publisher","first-page":"309","DOI":"10.1002\/mus.10443","volume":"28","author":"DG Kamper","year":"2003","unstructured":"Kamper, D. G., Harvey, R. L., Suresh, S. & Rymer, W. Z. Relative contributions of neural mechanisms versus muscle mechanics in promoting finger extension deficits following stroke. Muscle Nerve 28, 309\u2013318 (2003).","journal-title":"Muscle Nerve"},{"key":"98595_CR38","doi-asserted-by":"publisher","first-page":"1","DOI":"10.3389\/fpsyg.2018.02249","volume":"9","author":"RG Carson","year":"2018","unstructured":"Carson, R. G. & Rankin, M. L. Shaping the effects of associative brain stimulation by contractions of the opposite limb. Front. Psychol. 9, 1\u201312 (2018).","journal-title":"Front. Psychol."},{"key":"98595_CR39","doi-asserted-by":"publisher","first-page":"66","DOI":"10.1152\/jn.00383.2003","volume":"92","author":"K Stefan","year":"2004","unstructured":"Stefan, K., Wycislo, M. & Classen, J. Modulation of associative human motor cortical plasticity by attention. J. Neurophysiol. 92, 66\u201372 (2004).","journal-title":"J. Neurophysiol."},{"key":"98595_CR40","doi-asserted-by":"publisher","first-page":"501","DOI":"10.1113\/jphysiol.1993.sp019912","volume":"471","author":"T Kujirai","year":"1993","unstructured":"Kujirai, T. et al. Corticocortical Inhibition in human motor cortex. J. Physiol. 471, 501\u2013519 (1993).","journal-title":"J. Physiol."},{"key":"98595_CR41","doi-asserted-by":"publisher","unstructured":"Martino Cinnera, A. et al. Upper limb assessment with inertial measurement units according to the international classification of functioning in stroke: a systematic review and correlation meta-analysis. Top. Stroke Rehabil.   31, 66\u201385. https:\/\/doi.org\/10.1080\/10749357.2023.2197278   (2024).","DOI":"10.1080\/10749357.2023.2197278"},{"key":"98595_CR42","doi-asserted-by":"publisher","first-page":"13","DOI":"10.2340\/1650197771331","volume":"7","author":"AR Fugl-Meyer","year":"1975","unstructured":"Fugl-Meyer, A. R., J\u00e4\u00e4sk\u00f6, L., Leyman, I., Olsson, S. & Steglind, S. The post-stroke hemiplegic patient. 1. a method for evaluation of physical performance. Scand. J. Rehabil Med. 7, 13\u201331 (1975).","journal-title":"Scand. J. Rehabil Med."},{"key":"98595_CR43","doi-asserted-by":"publisher","first-page":"215","DOI":"10.2519\/jospt.1992.16.5.215","volume":"16","author":"GF Hamilton","year":"1992","unstructured":"Hamilton, G. F., McDonald, C. & Chenier, T. C. Measurement of grip strength: validity and reliability of the sphygmomanometer and Jamar grip dynamometer. J. Orthop. Sports Phys. Therapy 16, 215\u2013219 (1992).","journal-title":"J. Orthop. Sports Phys. Therapy"},{"key":"98595_CR44","doi-asserted-by":"publisher","first-page":"1713","DOI":"10.1007\/s10072-016-2650-6","volume":"37","author":"M Mancuso","year":"2016","unstructured":"Mancuso, M. et al. Italian normative data for a stroke specific cognitive screening tool: the Oxford cognitive screen (OCS). Neurol. Sci. 37, 1713\u20131721 (2016).","journal-title":"Neurol. Sci."},{"key":"98595_CR45","doi-asserted-by":"publisher","first-page":"615","DOI":"10.1007\/s00221-007-0960-x","volume":"181","author":"MV Sale","year":"2007","unstructured":"Sale, M. V., Ridding, M. C. & Nordstrom, M. A. Factors influencing the magnitude and reproducibility of corticomotor excitability changes induced by paired associative stimulation. Exp. Brain Res. 181, 615\u2013626 (2007).","journal-title":"Exp. Brain Res."},{"key":"98595_CR46","unstructured":"The Jamovi Project. Jamovi (version 2.6) [Computer Software]  (2025). https:\/\/www.jamovi.org."},{"key":"98595_CR47","unstructured":"R Core Team. R: A language and Environment for Statistical Computing    (2019). http:\/\/www.r-project.org\/."},{"key":"98595_CR48","doi-asserted-by":"publisher","unstructured":"Craighero, L. Motor resonance: neurophysiological origin, functional role, and contribution of the motivational, moral, and social aspects of action. In The Routledge Handbook of Embodied Cognition 442\u2013451 (Routledge, 2024). https:\/\/doi.org\/10.4324\/9781003322511-46.","DOI":"10.4324\/9781003322511-46"},{"key":"98595_CR49","doi-asserted-by":"publisher","first-page":"213","DOI":"10.1016\/j.conb.2005.03.013","volume":"15","author":"L Fadiga","year":"2005","unstructured":"Fadiga, L., Craighero, L. & Olivier, E. Human motor cortex excitability during the perception of others\u2019 action. Curr. Opin. Neurobiol. 15, 213\u2013218 (2005).","journal-title":"Curr. Opin. Neurobiol."},{"key":"98595_CR50","doi-asserted-by":"publisher","first-page":"655","DOI":"10.1152\/physrev.00009.2013","volume":"94","author":"G Rizzolatti","year":"2014","unstructured":"Rizzolatti, G., Cattaneo, L., Fabbri-Destro, M. & Rozzi, S. Cortical mechanisms underlying the organization of goal-directed actions and mirror neuron-based action Understanding. Physiol. Rev. 94, 655\u2013706 (2014).","journal-title":"Physiol. Rev."},{"key":"98595_CR51","doi-asserted-by":"publisher","first-page":"180","DOI":"10.1016\/j.cortex.2015.11.009","volume":"75","author":"TT de Beukelaar","year":"2016","unstructured":"de Beukelaar, T. T., Alaerts, K., Swinnen, S. P. & Wenderoth, N. Motor facilitation during action observation: the role of M1 and PMv in Grasp predictions. Cortex 75, 180\u2013192 (2016).","journal-title":"Cortex"},{"key":"98595_CR52","doi-asserted-by":"publisher","first-page":"2352","DOI":"10.1162\/jocn.2010.21590","volume":"23","author":"C Catmur","year":"2011","unstructured":"Catmur, C., Mars, R. B., Rushworth, M. F. & Heyes, C. Making mirrors: premotor cortex stimulation enhances mirror and counter-mirror motor facilitation. J. Cogn. Neurosci. 23, 2352\u20132362 (2011).","journal-title":"J. Cogn. Neurosci."},{"key":"98595_CR53","doi-asserted-by":"publisher","first-page":"211","DOI":"10.1016\/j.brs.2024.02.011","volume":"17","author":"E Chiappini","year":"2024","unstructured":"Chiappini, E. et al. Driving hebbian plasticity over ventral premotor-motor projections transiently enhances motor resonance. Brain Stimul. 17, 211\u2013220 (2024).","journal-title":"Brain Stimul"},{"key":"98595_CR54","doi-asserted-by":"publisher","first-page":"3757","DOI":"10.1162\/jocn_a_00051","volume":"23","author":"G Cantarero","year":"2011","unstructured":"Cantarero, G. et al. Disrupting the ventral premotor cortex interferes with the contribution of action observation to Use-dependent plasticity. J. Cogn. Neurosci. 23, 3757\u20133766 (2011).","journal-title":"J. Cogn. Neurosci."},{"key":"98595_CR55","doi-asserted-by":"publisher","first-page":"419","DOI":"10.1007\/s00429-005-0039-z","volume":"210","author":"G Rizzolatti","year":"2005","unstructured":"Rizzolatti, G. The mirror neuron system and its function in humans. Anat. Embryol. (Berl). 210, 419\u2013421 (2005).","journal-title":"Anat. Embryol. (Berl)"},{"key":"98595_CR56","doi-asserted-by":"crossref","unstructured":"Alaerts, K., Swinnen, S. P. & Wenderoth, N. Is the human primary motor cortex activated by muscular or direction-dependent features of observed movements? Cortex 45, 1148\u20131155 (2009).","DOI":"10.1016\/j.cortex.2008.10.005"},{"key":"98595_CR57","doi-asserted-by":"publisher","first-page":"e2404925121","DOI":"10.1073\/pnas.2404925121","volume":"121","author":"S Turrini","year":"2024","unstructured":"Turrini, S. et al. Spike-timing-dependent plasticity induction reveals dissociable supplementary- and premotor-motor pathways to automatic imitation. Proc. Natl. Acad. Sci. U S A. 121, e2404925121 (2024).","journal-title":"Proc. Natl. Acad. Sci. U S A"},{"key":"98595_CR58","doi-asserted-by":"publisher","first-page":"173","DOI":"10.1016\/j.cortex.2020.01.008","volume":"126","author":"O Maddaluno","year":"2020","unstructured":"Maddaluno, O., Guidali, G., Zazio, A., Miniussi, C. & Bolognini, N. Touch anticipation mediates cross-modal hebbian plasticity in the primary somatosensory cortex. Cortex 126, 173\u2013181 (2020).","journal-title":"Cortex"},{"key":"98595_CR59","doi-asserted-by":"publisher","first-page":"1109","DOI":"10.1038\/nn.2182","volume":"11","author":"SM Aglioti","year":"2008","unstructured":"Aglioti, S. M., Cesari, P., Romani, M. & Urgesi, C. Action anticipation and motor resonance in elite basketball players. Nat. Neurosci. 11, 1109\u20131116 (2008).","journal-title":"Nat. Neurosci."},{"key":"98595_CR60","doi-asserted-by":"crossref","unstructured":"Qin, C. et al. Predictability alters information flow during action observation in human electrocorticographic activity. Cell. Rep. 42, 456 (2023).","DOI":"10.1016\/j.celrep.2023.113432"},{"key":"98595_CR61","doi-asserted-by":"publisher","first-page":"2511","DOI":"10.1093\/cercor\/bhp292","volume":"20","author":"C Urgesi","year":"2010","unstructured":"Urgesi, C. et al. Simulating the future of actions in the human corticospinal system. Cereb. Cortex. 20, 2511\u20132521 (2010).","journal-title":"Cereb. Cortex"},{"key":"98595_CR62","doi-asserted-by":"publisher","first-page":"710","DOI":"10.1093\/cercor\/bhr157","volume":"22","author":"A Cavallo","year":"2012","unstructured":"Cavallo, A., Becchio, C., Sartori, L., Bucchioni, G. & Castiello, U. Grasping with tools: corticospinal excitability reflects observed hand movements. Cereb. Cortex. 22, 710\u2013716 (2012).","journal-title":"Cereb. Cortex"},{"key":"98595_CR63","doi-asserted-by":"crossref","unstructured":"Gangitano, M.,  Mottaghy, F. M. & Pascual-leone, A. Phase-specific modulation of cortical motor output during movement observation. Cortex 12, 1489\u20131492 (2001).","DOI":"10.1097\/00001756-200105250-00038"},{"key":"98595_CR64","doi-asserted-by":"publisher","first-page":"3513","DOI":"10.1016\/j.neuropsychologia.2010.07.037","volume":"48","author":"G Koch","year":"2010","unstructured":"Koch, G. et al. Resonance of cortico-cortical connections of the motor system with the observation of goal directed grasping movements. Neuropsychologia 48, 3513\u20133520 (2010).","journal-title":"Neuropsychologia"},{"key":"98595_CR65","doi-asserted-by":"publisher","first-page":"43","DOI":"10.1016\/j.cortex.2016.09.002","volume":"84","author":"KL Bunday","year":"2016","unstructured":"Bunday, K. L., Lemon, R. N., Kilner, J. M., Davare, M. & Orban, G. A. Grasp-specific motor resonance is influenced by the visibility of the observed actor. Cortex 84, 43\u201354 (2016).","journal-title":"Cortex"},{"key":"98595_CR66","doi-asserted-by":"publisher","first-page":"74","DOI":"10.1016\/j.neuroimage.2016.03.060","volume":"134","author":"L Amoruso","year":"2016","unstructured":"Amoruso, L. & Urgesi, C. Contextual modulation of motor resonance during the observation of everyday actions. Neuroimage 134, 74\u201384 (2016).","journal-title":"Neuroimage"},{"key":"98595_CR67","doi-asserted-by":"crossref","unstructured":"Anson, J. G., Hasegawa, Y., Kasai, T., Latash, M. L. & Yahagi, S. EMG discharge patterns during human grip movement are Task-Dependent and not modulated by muscle contraction modes: a transcranial magnetic stimulation (TMS) study. Brain Res. 2002, 934 (2002). www.elsevier.com\/locate\/bres.","DOI":"10.1016\/S0006-8993(02)02368-5"},{"key":"98595_CR68","doi-asserted-by":"publisher","first-page":"1513","DOI":"10.1111\/j.1460-9568.2005.04336.x","volume":"22","author":"M Montagna","year":"2005","unstructured":"Montagna, M., Cerri, G., Borroni, P. & Baldissera, F. Excitability changes in human corticospinal projections to muscles moving hand and fingers while viewing a reaching and grasping action. Eur. J. Neurosci. 22, 1513\u20131520 (2005).","journal-title":"Eur. J. Neurosci."},{"key":"98595_CR69","unstructured":"Neumann, D. D. Kinesology of Musculoskeletal System  465\u2013688 (Mosby Inc, 2010)."},{"key":"98595_CR70","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1371\/journal.pone.0177457","volume":"12","author":"G Puglisi","year":"2017","unstructured":"Puglisi, G. et al. The role of attention in human motor resonance. PLoS One 12, 1\u201321 (2017).","journal-title":"PLoS One"},{"key":"98595_CR71","doi-asserted-by":"publisher","first-page":"235","DOI":"10.1007\/s00221-010-2358-4","volume":"205","author":"S Schuch","year":"2010","unstructured":"Schuch, S., Bayliss, A. P., Klein, C. & Tipper, S. P. Attention modulates motor system activation during action observation: evidence for inhibitory rebound. Exp. Brain Res. 205, 235\u2013249 (2010).","journal-title":"Exp. Brain Res."},{"key":"98595_CR72","doi-asserted-by":"publisher","first-page":"1","DOI":"10.3389\/fnhum.2013.00823","volume":"7","author":"RG Carson","year":"2013","unstructured":"Carson, R. G. & Kennedy, N. C. Modulation of human corticospinal excitability by paired associative stimulation. Front. Hum. Neurosci. 7, 1\u201328 (2013).","journal-title":"Front. Hum. Neurosci."},{"key":"98595_CR73","doi-asserted-by":"publisher","first-page":"3929","DOI":"10.1523\/JNEUROSCI.1870-17.2018","volume":"38","author":"SA Aguiar","year":"2018","unstructured":"Aguiar, S. A. & Baker, S. N. Convergent spinal circuits facilitating human wrist flexors. J. Neurosci. 38, 3929\u20133938 (2018).","journal-title":"J. Neurosci."},{"key":"98595_CR74","doi-asserted-by":"crossref","unstructured":"Pierrot-Deseilligny, E. & Burke, D. The Circuitry of the Human Spinal Cord: Spinal and Corticospinal Mechanisms of Movement (Cambridge University Press, 2012).","DOI":"10.1017\/CBO9781139026727"},{"key":"98595_CR75","doi-asserted-by":"publisher","first-page":"1","DOI":"10.3389\/fnins.2019.00841","volume":"13","author":"L Minkova","year":"2019","unstructured":"Minkova, L. et al. Determinants of inter-individual variability in corticomotor excitability induced by paired associative stimulation. Front. Neurosci. 13, 1\u201314 (2019).","journal-title":"Front. Neurosci."},{"key":"98595_CR76","doi-asserted-by":"crossref","unstructured":"Strube, W., Bunse, T., Malchow, B. & Hasan, A. Efficacy and interindividual variability in motor-cortex plasticity following anodal tDCS and paired-associative stimulation. Neural Plast.  (2015).","DOI":"10.1155\/2015\/530423"},{"key":"98595_CR77","first-page":"183","volume":"36","author":"JA Palmer","year":"2018","unstructured":"Palmer, J. A., Wolf, S. L. & Borich, M. R. Paired associative stimulation modulates corticomotor excitability in chronic stroke: a preliminary investigation. Restor. Neurol. Neurosci. 36, 183\u2013194 (2018).","journal-title":"Restor. Neurol. Neurosci."},{"key":"98595_CR78","first-page":"281","volume":"22","author":"G Kwakkel","year":"2004","unstructured":"Kwakkel, G., Kollen, B. & Lindeman, E. Understanding the pattern of functional recovery after stroke: facts and theories. Restor. Neurol. Neurosci. 22, 281\u2013299 (2004).","journal-title":"Restor. Neurol. Neurosci."},{"key":"98595_CR79","doi-asserted-by":"crossref","unstructured":"Bernard-Espina, J., Beraneck, M., Maier, M. A. & Tagliabue, M. Multisensory integration in stroke patients: a theoretical approach to reinterpret Upper-Limb proprioceptive deficits and visual compensation. Front. Neurosci. 15, 74 (2021).","DOI":"10.3389\/fnins.2021.646698"},{"key":"98595_CR80","doi-asserted-by":"crossref","unstructured":"Ma, L., Marshall, P. J. & Wright, W. G. The impact of external and internal focus of attention on visual dependence and EEG alpha oscillations during postural control. J. Neuroeng. Rehabil 19, 145 (2022).","DOI":"10.1186\/s12984-022-01059-7"},{"key":"98595_CR81","doi-asserted-by":"publisher","first-page":"607","DOI":"10.1111\/j.1469-7793.1998.607bn.x","volume":"509","author":"R Hanajima","year":"1998","unstructured":"Hanajima, R. et al. Paired-pulse magnetic stimulation of the human motor cortex: differences among I waves. J. Physiol. 509, 607\u2013618 (1998).","journal-title":"J. Physiol."},{"key":"98595_CR82","doi-asserted-by":"crossref","unstructured":"Butefisch, C. M. Mechanisms of use-dependent plasticity in the human motor cortex. Proc.  Natl. Acad. Sci 97, 3661\u20133665 (2000).","DOI":"10.1073\/pnas.050350297"},{"key":"98595_CR83","doi-asserted-by":"publisher","first-page":"2586","DOI":"10.1093\/brain\/awg273","volume":"126","author":"A Quartarone","year":"2003","unstructured":"Quartarone, A. et al. Abnormal associative plasticity of the human motor cortex in Writer\u2019s cramp. Brain 126, 2586\u20132596 (2003).","journal-title":"Brain"},{"key":"98595_CR84","doi-asserted-by":"publisher","first-page":"363","DOI":"10.1016\/j.jocn.2021.06.028","volume":"90","author":"YF Sui","year":"2021","unstructured":"Sui, Y. F., Tong, L. Q., Zhang, X. Y., Song, Z. H. & Guo, T. C. Effects of paired associated stimulation with different stimulation position on motor cortex excitability and upper limb motor function in patients with cerebral infarction. J. Clin. Neurosci. 90, 363\u2013369 (2021).","journal-title":"J. Clin. Neurosci."},{"key":"98595_CR85","doi-asserted-by":"crossref","unstructured":"Liang, S. et al. Repetitive peripheral magnetic stimulation combined with transcranial magnetic stimulation in rehabilitation of upper extremity hemiparesis following stroke: a pilot study. J. Rehabil Med. 56, 45 (2024).","DOI":"10.2340\/jrm.v56.19449"},{"key":"98595_CR86","doi-asserted-by":"publisher","first-page":"341","DOI":"10.1016\/j.neubiorev.2011.07.004","volume":"36","author":"P Molenberghs","year":"2012","unstructured":"Molenberghs, P., Cunnington, R. & Mattingley, J. B. Brain regions with mirror properties: a meta-analysis of 125 human fMRI studies. Neurosci. Biobehav Rev. 36, 341\u2013349 (2012).","journal-title":"Neurosci. Biobehav Rev."},{"key":"98595_CR87","doi-asserted-by":"publisher","first-page":"16","DOI":"10.1016\/j.cortex.2022.11.007","volume":"159","author":"L Craighero","year":"2023","unstructured":"Craighero, L., Mele, S., Gaifas, V., Bonaguri, E. & Straudi, S. Evidence of motor resonance in stroke patients with severe upper limb function impairments. Cortex 159, 16\u201325 (2023).","journal-title":"Cortex"},{"key":"98595_CR88","first-page":"571","volume":"34","author":"N Bolognini","year":"2016","unstructured":"Bolognini, N., Russo, C. & Edwards, D. J. The sensory side of post-stroke motor rehabilitation. Restor. Neurol. Neurosci. 34, 571\u2013586 (2016).","journal-title":"Restor. Neurol. Neurosci."},{"key":"98595_CR89","doi-asserted-by":"publisher","unstructured":"Dimyan, M. A. & Cohen, L. G. Neuroplasticity in the context of motor rehabilitation after stroke. Nat. Rev. Neurol. 7, 76\u201385. https:\/\/doi.org\/10.1038\/nrneurol.2010.200   (2011).","DOI":"10.1038\/nrneurol.2010.200"}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-98595-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-98595-8","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-98595-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,5,1]],"date-time":"2025-05-01T17:03:01Z","timestamp":1746118981000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-98595-8"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,5,1]]},"references-count":89,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2025,12]]}},"alternative-id":["98595"],"URL":"https:\/\/doi.org\/10.1038\/s41598-025-98595-8","relation":{"has-preprint":[{"id-type":"doi","id":"10.1101\/2024.08.27.24312576","asserted-by":"object"}]},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,5,1]]},"assertion":[{"value":"13 December 2024","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"14 April 2025","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"1 May 2025","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"15313"}}